The Influence of Fire on Saltatoria Diversity in Coastal Habitats Near Pangani,Tanzania (East Africa)

ECOTROPICA 11: 53–61, 2005 © Society for Tropical Ecology

THE INFLUENCE OF FIRE ON SALTATORIA DIVERSITY IN COASTAL HABITATS NEAR PANGANI,TANZANIA (EAST AFRICA)

Claudia Hemp

Dept. Animal Ecology II, University of Bayreuth, 95440 Bayreuth

Abstract. In burnt coastal grasslands, fire-disturbed forest remnants, and undisturbed coastal forests of the Pangani area (Tanzania) 61 Saltatoria species were recorded. Comparing open-land habitats with forest and forest edge habitats, diversity rises in open-land habitats, as nearly 60% of the recorded species were found within grasslands. However, the share of endemic and near-endemic species decreases, as only 5% of the species occurring in open-land habitats were endemics, 14% are distributed throughout East Africa, and the majority (81%) are widespread forms. Forest and forest edge habitats, on the other hand, had 60% endemic species, 32% species restricted to East Africa, and only 8% widespread Saltatoria. The majority of the coastal endemics are highly endangered by the ongoing destruction of indigenous forest. Tangana asymmetrica was one of the few coastal endemics that also occurred in fire-influenced forest remnants, while most other forest species were exclusively found in closed undisturbed forest communities of the Gendagenda forest reserve. The new record of Parepistaurus pygmaeus in the Gendagenda reserve offers an explanation of the hitherto disjunct distribution of this flightless species. Accepted 15 June 2005. Key words: Biodiversity, conservation, endemism, fire influence, indigenous tropical lowland forest, Orthoptera, Saltatoria, Tanzania, tropical coastal grasslands.

INTRODUCTION lived in the area, mostly farmers, cultivating the fields intensively with bananas, maize, sweet potatoes, ma- The forests along the Kenyan and Tanzanian coast are nihot etc. (Koponen 1994). the remains of a once widespread forest cover. Today Away from the main centers of cultivation around this forest has largely gone, leaving around 250 patches of forest, most of which are less than 500 ha in size Pangani the coastal area was probably little populated. (Burgess & Clarke 2000). The coastal forests, together Most of the coast and hinterland was densely forested. with the adjacent forests of the Eastern Arc Mountains, Clearing by fires was practised from the 19th century are hotspots of endemism due to a constant climate onwards, reducing the forests especially along the and long isolation from the Central African forests coastal line, mainly for establishing sisal plantations (Rodgers & Homewood 1982). A stable long-term and crop production. climate in combination with high rainfall in tropical About 20 km inland from Pangani an elevation forest areas are usually associated with a high species step of about 50–100 m is apparent. Here at about diversity (Fjeldså & Lovett 1997). A loss of these for- 300 m a.s.l. stretches dense coastal forest belonging ests would result in an enormous loss of biodiversity to the Gendagenda reserve. However, in the past 10 (Newmark 2002). years even here the practice of setting fires to reduce In the vicinity of the studied plots lies the coastal tree cover in favor of grassland was observed. It must town Pangani, which had its greatest importance in be feared that also this patch of indigenous lowland the 19th century. Pangani was the starting and desti- forest will disappear in the near future. nation of caravans coming from or heading for the The aim of this study was to compare the Salta- hinterland, trading with ivory and later with slaves toria composition of intact coastal forest within the (Glassman 1995). In 1880 about 100 000 people Gendagenda reserve with fire-influenced patches of forest and grasslands resulting from intensive burning over the last decades near Mwera village, south of e-mail: [email protected] Pangani (Fig. 1). 53 HEMP

FIG. 1. Map of East Africa. Black triangles: study plots 1: fire-maintained grasslands and fire-influenced coastal forests near Mwera. 2: Gen- dagenda forest reserve.

RESEARCH AREA some parts of which are already disturbed by infre- quent fires (Fig. 2). The coastal area of Tanzania receives its precipitation Burnt grasslands were investigated directly at the from monsoonal rainfalls from the Indian Ocean co- coast near Pangani, about 20 km southward, next to inciding with the northward and southward passages the small village of Mwera (UTM zone 37 04/99/369 in March to May (long rains with most precipitation) east, 93/94/462 south) (Figs. 1, 2). Plant species com- and October to December (short rains). The district mon in coastal savanna grasslands were the grasses city Tanga receives a total of about 1000 mm per Imperata cylindrica, Digitaria milanjiana, Echinochloa year. haploclada, Setaria incrassata, Panicum infestum and The study area is characterized by huge sisal plan- the herb Kohautia coccinea (A. Hemp unpubl. data). tations about 5–10 km inland, and wide patches of The vegetation cover was 80–100% with a height of burnt grasslands intermingled with remnants of for- about 1m. These grasslands are set on fire nearly every est (East African coastal scrub forest, Burgess & Clarke year before the long rains in March. 2000). Adjacent to the sisal plantations the indige- Remnants of forest are intermingled with the grass- nous forest (East African coastal dry forest, Burgess lands. After frequent fires the fire-tolerant palmtree & Clarke 2000) of the Gendagenda reserve is situated, Hyphaene compressa sprouts. Well adapted to frequent 54 EAST AFRICAN SALTATORIA fires, the South American tree Leucaena latisiliqua becomes dominant in the first years after a fire, form- ing nearly monospecific stands. In the regeneration stage after about 10 to 15 years following a fire indigenous trees like Kigelia africana, Balanites wilsoniana, Lannea schweinfurthii, Xanthoxylum cf. holtzianum, Guettarda speciosa, and Sclerocarya birrea form the tree layer, and shrubs such as Psychotria lauracea, Grewia forbesii, Allophylus rubifolius, Harrisonia abyssinica, Vangueria infausta, Flueggea virosa, Pemphis acidula and Polysphae- ria multiflora are found in the undergrowth (A. Hemp unpubl. data). The canopy is usually 10 m high. All investigated forests remnants along the coast in the FIG. 2. Burnt grassland at the coast near the village area belong to regeneration stages after fire. Some cen- Mwera on the Indian Ocean. In the background the tral parts of these remnants might be more or less un- fire-tolerant palmtree Hyphaene compressa. disturbed by fires, although the patches are mostly not bigger than a few square meters (equivalent to the forest community “coastal scrub forest” in Burgess & Clarke 2000). closed forest. In the closed forest, whether the spe- Areas of undisturbed indigenous lowland forest cies occurred on the forest floor or in the bush or tree mixed with fire-influenced forests were investigated layer was registered. In grasslands, Saltatoria species between Kabuko and Mwera at 350 m a.s.l. (Gen- were identified by sight by walking repeatedly on par- dagenda forest reserve, Fig. 3). This forest type, allel tracks through the plots at distances of about which belongs to legume-dominated dry forest, after 1–1.5 m. Smaller species and those difficult to iden- Burgess & Clarke (2000), was characterized by trees tify were caught by net-sweeping. The abundance of and shrubs such as Brachystegia spiciformis, Milicia gomphocerines was estimated by the male courtship excelsa, Strychnos scheffleri, Hugonia castaneifolia, Po- song. In forest habitats all Saltatoria were caught, as lyaltia stuhlmannii, Kraussia kirkii, Polysphaeria parvi- the density of individuals was generally low. Bushes folia, Psychotria punctata and Pavetta stenosepala (A. and small trees were checked by net-sweeping. The Hemp, unpubl. data). The mean canopy height was check of one plot lasted in general more than 1 hour, 25 m. in forest habitats sometimes more than 2 hours. Near

METHODS Saltatoria species composition was recorded for plots of burnt coastal grasslands, for several patches of forest remnants and an adjacent larger forest patch in a regeneration stage (indicated with “1” in Fig. 1), and two plots of dense indigenous forest near the Kabuko- Mwera road (Gendagenda forest reserve, indicated with “2” in Figure 1). All plots were monitored several times (February 1999, January 2000, January and February 2001, December 2004). Four plots were chosen in homogenous grasslands, usually an area of 30 x 30 m. The forest plots had sizes of about 100 x 100 m (two plots within the Gendagenda forest reserve, one plot in regeneration stage, three fire-influenced plots). FIG. 3. Coastal forest area at 300 m a.s.l. between Ka- In the forest plots it was noted in which micro- buko and Mwera. In the background the Gendagenda habitat each species occurred. A differentiation was Hill. In the foreground fire impact along the road side made between forest edge (clearing or road side) and vegetation. 55 HEMP

TABLE 1. List of Saltatoria. Data on habitat and distribution. Gr: grassland species, Fe: forest edge species, F: forest species, Dis: distribution, EA: species restricted to East Africa, C: species restricted to coastal habitats (coastal endemics), remaining species: widespread; Loc: 1: coastal forest remnants and grasslands near Mwera; 2: Gendagenda forest reserve (as shown on Fig. 1).

Species/Family Habitat Gr Fe F Dis loc Family Tetrigidae Paratettix africanus Bolivar, 1908 Open patches in humid grasslands x 1 Family Eumastacidae Plagiotriptus hippiscus (Gerstäcker, 1869) Forest and forest edge x EA 1,2 Euschmidtia sansibarica Karsch, 1889 Bushes at forest edges x C 1,2 Family Lentulidae Usambilla sp.* Undergrowth of coastal forest x C 2 Family Acrididae Subfamily Hemiacridinae Leptacris monteiroi monteiroi (I. Bolivar, 1890) Burnt grasslands x 1 Meruana usambarica Karsch, 1896 Burnt grasslands x EA 1 Oraistes luridus Karsch, 1896 Shrub layer of coastal forest x EA 2 Paraspathosternum pedestris (Miller, 1929) Undergrowth of coastal forest x C 2 Subfamily Oxyinae Oxya hyla hyla Serville, 1831 Humid places of burnt grasslands x 1 Subfamily Coptacridinae Parepistaurus pygmaeus (Karny, 1909) Undergrowth of coastal forest x C 2 Subfamily Eyprepocnemidinae Cataloipus oberthuri oberthuri (I. Bolivar, 1890) Burnt grasslands x 1 Eyprepocnemis plorans ibandana (Gig.-Tos, Burnt grasslands; 1907) attracted to light at night x 1 Heteracris coerulipes (Sjöstedt, 1909) Undergrowth at forest edges x C 2 Metaxymecus gracilipes (Brancsik, 1895) Burnt grasslands and forest edges x 1 Taramassus cunctator cunctator (Karsch, 1900) Burnt grasslands x EA 1 Paraprocticus forchhammeri (Johnsen, 1974) Forest floor and forest edge x C 2 Subfamily Catantopinae Abisares viridipennis viridipennis (Bur., 1838) Forests edges x 2 Catantops momboensis momboensis Sjöstedt, Burnt grasslands, 1931 bushland and forest edges x EA 1 Diabolocatantops axillaris axillaris (Bur., Open sand and coastal grasslands 1838) sparse in vegetation x 1 Eupropacris vana (Karsch, 1896) On bushes in the understory of coastal forest x EA 2 Eupropacris pompalis (Karsch, 1896) Herb vegetation along forest edges x C 1 Tangana asymmetrica Ramme, 1929 Litter of coastal forest x C 1,2 Phaeocatantops decoratus (Gerstäcker, 1869) Coastal bush and forest edges x EA 1,2 Pseudophialosphera tectifera (Ramme, 1929) Undergrowth of coastal forest x C 2 Subfamily Cyrtacanthacridinae Ornithacris cyanea (Stoll, 1813) Burnt grasslands x 1 Subfamily Tropidopolinae Tristia marginicosta Karsch, 1896 Burnt grasslands x 1 Subfamily Oedipodinae Ailopus thalassinus (Fabricius, 1781) Disturbed places x 1 Gastrimargus africanus africanus (Saussure, 1888) Burnt grasslands, disturbed places x 1 Heteropternis thoracica (Walker, 1870) Burnt grasslands x 1

56 EAST AFRICAN SALTATORIA

Table 1 continued

Species/Family Habitat Gr Fe F Dis loc

Humbe tenuicornis (Schaum, 1853) Burnt grasslands x 1 Jasomenia sansibara (Karsch, 1896) Burnt grasslands, especially in more humid places; attracted to light at night x 1 Morphacris fasciata (Thunberg, 1815) Disturbed places x 1 Pternoscirtes pallidus (Walker, 1870) Open sand at the shore x 1 Subfamily Acridinae Acrida bicolor (Thunberg, 1815) Burnt grasslands, disturbed places x 1 Acrida sulphuripennis (Gerstäcker, 1869) Burnt grasslands, disturbed places x 1 Afrophlaeoba usambarica (Ramme, 1929) Undergrowth of coastal forest x C 2 Comacris semicarinatus (Gerstäcker, 1869) Burnt grasslands x EA 1 Gymnobothrus temporalis flexuosus Semi-shade on paths (Schulthess, 1898) and forest edges x EA 1 Lobopoma mitchelli Popov & Fishpool, 1992 Burnt grasslands x C 1 Machaeridia conspersa I. Bolivar, 1889 Burnt grasslands x 1 Orthochtha dasycnemis (Gerstäcker 1869) Burnt grasslands x 1 Orthochtha dimorpha Miller, 1929 Burnt grasslands x EA 1 Subfamily Gomphocerinae Minihippus keyi (Uvarov, 1941) Burnt grasslands x C 1 Brachycrotaphus sjostedti Uvarov, 1932 Burnt grasslands x EA 1 Mesopsis abbreviatus (Beauvois, 1806) Burnt grasslands x 1 Mesopsis laticornis (Krauss, 1877) Burnt grasslands x 1 Pnorisa sp. Burnt grasslands x 1 Pnorisa squalus squalus (Stål, 1860) Burnt grasslands x 1 Family Tettigoniidae Subfamily Phaneropterinae Catoptropteryx aurita Huxley, 1970 Attracted to light from remnants of coastal forest x EA 1,2 Eurycorypha sp.* Attracted to light from remnants of coastal forest x C 1,2 Horatosphaga heteromorpha (Karsch, 1888) Burnt grasslands; attracted to light at night x EA 1 Dioncomena ornata Brunner v. Wattenwyl, 1878 Shrub layer of coastal forest x C 2 Ducetia biramosa (Karsch, 1888) Forest edge in coastal forest x C 1,2 Tylopsis rubrescens Kirby, 1900 Burnt grasslands x 1 Tylopsis irregularis Karsch, 1893 Burnt grasslands x 1 Subfamily Mecopodinae Gymnoscirtus unguiculatus (Karsch, 1888) Undergrowth of coastal forest x C 2 Subfamily Conocephalinae Agraecia sansibara Redtenbacher, 1891 Attracted to light near forest x C 1,2 Conocephalus (C.) conocephalus (Linné, 1767) Burnt grasslands x 1 Megalotheca longiceps (Peringuey, 1916) Burnt grasslands x 1 Subfamily Pseudophyllinae Acauloplax exigua Karsch, 1891 Tree layer of coastal forest x EA 2 Subfamily Hetrodinae Enyaliopsis bloyeti (Lucas, 1885) Bushes along forest edges x C 1

* probably new species, thus it is assumed that these Saltatoria are coastal endemics 57 HEMP

TABLE 2. Habitat and distribution pattern of Saltatoria species on Pangani coast. WS: widespread species (whole Africa, tropical Africa), EA: species confined to East Africa, C: coastal endemics, with share of the total number in %, Total: total species number.

Habitat WS % EA % C % Total Grasslands 29 81% 5 14% 2 5% 36 Forest edge 2 15% 4 31% 7 54% 13 Forest 0 – 4 33% 8 67% 12 Totals 32 52% 12 20% 17 28% 61

the forest remnants along the coast (indicated with perform courtship by rising up from high grasses with “1” in Fig. 1) light traps were used to register tetti- a flying sound similar to that of the European acridid goniids. Psophus stridulus (Linné, 1758). In living specimens the color of the alae is bright orange, fading after the RESULTS insects have been preserved. A total of 48 Acridoidea (with Tetrigidae) and 13 The majority of species occurring along forest Tettigonioidea species were recorded from the area edges are endemics of coastal habitats (54%) or are (Table 1). restricted to East Africa (31%) (Table 2). In the grasslands, 36 species were found, most of Heteracris coerulipes is conspicuous because of them (29 species) widespread in Africa (Table 1). Five its colorful alae (Fig. 4). This eyprepocnemidine is of the grassland species are restricted to East Africa known from the forests of coastal Tanzania and the and only two were coastal forms. Eastern Arc Mountains (Grunshaw 1991). It was High frequencies of occurrence (80–100% pre- found inhabiting patches of grass along forest paths sence during all checks of the plots) were seen in the and grassy forest clearings in the forested area between species Orthochtha dasycnemis, Conocephalus conoce- Kabuko and Mwera. Individuals of this species were phalus, Ornithacris cyanea, Eyprepocnemis plorans and collected with red or blue alae. The hetrodine Enya- Comacris semicarinatus. Some of these species also re- liopsis bloyeti, restricted to coastal Tanzania (Glenn ached a high abundance. Especially Orthochtha dasyc- 1991), is a species of forest edge and also bushland. nemis and Conocephalus conocephalus had population During the day individuals of this species are hardly densities of up to 2 individuals per square meter. Co- obtained while during the evening and night hours macris semicarinatus were also very conspicuous. Males males, climbing small bushes and trees to a height of 1–2 m, can be located by their loud song. Afrophlae- oba usambarica populated grassy patches on forest clearings and along forest paths. This species was previously known only from the East Usambara Mts. (Jago 1983). Parepistaurus pygmaeus (Fig. 5) is a coptacridine presently known from the Usambara and Nguru Mts. of the Eastern Arc Range. The records from lowland forest are interesting since Green (1998) discussed its disjunct distribution and possible reasons for it (see Discussion). In closed forest 12 Saltatoria species were recorded. All of them were either endemics (67%) or restricted to East Africa (33%, Table 2). Eight species occurred only in undisturbed indigenous forest, FIG. 4. Heteracris coerulipes (adult male, Eyprepoc- four species both in the Gendagenda forest reserve and nemidinae) is an inhabitant of forest edges. the coastal forest patch in regeneration (Table 1). 58 EAST AFRICAN SALTATORIA

by Ramme (1929) from the Tanga region and no additional information about this species has been hitherto published. Individuals of this species were found in closed forest of the Gendagenda reserve. Among the litter of the forest floor these flightless yellowish-brownish grasshoppers are well camouflaged. In escaping they showed similar habits as other flightless species e.g., Tangana asymmetrica (Fig. 7). In great jumps they tried to escape, landing among the yellowish-brownish leaves of the forest floor and sitting motionless till another disturbance induced them to move on.

DISCUSSION FIG. 5. Parepistaurus pygmaeus (Coptacridinae) was The Saltatoria diversity of the studied grasslands is previously known from the Usambara and Nguru Mts. higher compared with the species inventory of forests. However the percentage of coastal endemics is by far higher in forest (67%) than in grasslands (5%), where the majority of Saltatoria are widespread forms (81%, Records of Saltatoria species mentioned below Table 2, Fig. 8). Similar observations were made either extend our knowledge (since they were not by Cleary & Genner (2004) for butterflies and dragon- known from coastal forest), or, for some of them, flies in Borneo. The quality of species changes as en- description data only are available, mostly without any demic species disappear in favor of generalist spe- information on habitat or ecology. cies in fire-influenced rain forest communities. The- Catoptropteryx aurita is the only species of the se authors also found that a recolonization depends genus not occurring in West Africa. Nothing has been largely on the geographical distance between undis- published on the biology of this or other species of turbed habitats. As Saltatoria species are generally this genus. C.aurita is easily attracted to light during bound to certain microclimatic conditions, a recolo- the evening hours. This canopy inhabitant occurred nization of forest Saltatoria requiring high air humi- both in forest remnants at the coast near Mwera and dity seems nearly impossible under present-day con- in indiginous forest of the Gendagenda forest reserve. ditions, as forest patches are too far apart and are se- On Kilimanjaro it is reported also in the highly anthro- parated by grasslands and agricultural land with low pogenic Chagga home gardens, where it preferred the air humidity. Especially the forest remnants investi- tree Margaritaria discoidea (Hemp 2005). Agraecia sansibara was attracted to light at the coast near Mwera, probably from the surrounding patches of forest. It was known to be present only from Zanzibar (Redtenbacher 1891), this island being not far from the shore of the area investigated. It was also collected from bushes at night in the East Usam- bara Mts. (Hemp, unpubl. data). Gymnoscirtus unguiculatus is a lowland species known from localities in the Usambara Mts. (Karsch 1888, Sjöstedt 1909, Hemp 2002). This huge meco- podine occurred in closed forest between Kabuko and Mwera, where it was caught on the forest floor between litter. On the Zigi Trail (450 m a.s.l.) of the East Usambara Mts. it was obtained at night also from the forest floor. FIG. 6. The well camouflaged Pseudophialosphera tec- Pseudophialosphera tectifera (Fig. 6) is a catanto- tifera (adult female, Catantopinae) is an endemic of pine restricted to coastal forests. It was described coastal forest of Tanzania. 59 HEMP gated along the coast near Mwera, which are fire-in- ferent genus status refers solely to the asymmetric fluenced, are not close enough to intact forests to gua- form of the male genitalia in Tangana) seem to be rantee a recolonization by the whole array of forest capable of adapting to changing environmental con- species, especially flightless forms. This might explain ditions. Ixalidium sjostedti Kevan, 1950 on Mt. why, even in the relatively large patch of forest near Kilimanjaro and at Meru, and I. haematoscelis Ger- Mwera (in Fig. 1 indicated with 1) which had not stäcker, 1869 occurring in the Usambara Mts. and been affected by fire for at least 10–15 years and adjacent regions, for example, have obviously adapted which seemed to offer adequate habitat conditions to changing environmental conditions in the process again for forest species, only few forest species were of the ongoing destruction of indigenous forest. They noted there. Those species found in this forest were were found today to inhabit altitudinally higher areas, either forest edge species, with a wider ecological span, often forest edges (I. haematoscelis near Amani, E. which could have survived here (e.g., Plagiotriptus Usambara) and even open grassland communities (I. hippiscus, even occurring in degraded bushland, Eu- sjostedti, high altitude grasslands, see Hemp & Hemp schmidtia sansibara, Ducetia biramosa and Phaeoca- (2003)). Tangana asymmetrica seems to have the same tantops decoratus) or were fully winged, such as Agrae- flexibility as it is able to persist in landscapes where cia sansibara, Eurycorypha sp. and Catoptropteryx forest cover has been reduced to small patches. aurita. Tangana asymmetrica (Fig. 8) was the only As lowland forests have been fragmented by hu- flightless forest species which also occurred in the fire- man influence over the last centuries and even mil- influenced forest remnants at the coast near Mwera. lennia affecting especially the foothills of mountains Species of the genera Ixalidium and Tangana (the dif- ranges such as the Usambara, Pare or Uluguru Mts. (Burgess and Clarke 2000) species might have been fragmented in their occurrence as well. In this connec- tion the record of the flightless species Parepistaurus pygmaeus, previously known only from the Usambara and Nguru Mts. (Green 1998), provides an explanation for the hitherto assumed disjunct distribution in the Gendagenda forest reserve. Formerly this species clearly populated the whole area between the Usam- bara and Nguru Mts, but became extinct in the area between because of loss of adequate habitat. Since it has now been recorded in the Gendagenda reserve, lying somewhat intermediate and also indicating that this species is not restricted to submontane forest communities, it clearly shows that a changing environ- ment was responsible for the retreat of the species. Thus flightless species such as Parepistaurus pygmaeus clearly indicate that forest habitats formerly must have covered huge areas of coastal Tanzania, extending inland even to the ranges of the Eastern Arc complex such as the Nguru, Uluguru and West Usambara Mts., and which are replaced today, due to human influence, by habitat types common to large areas of Africa. Hand in hand with the loss of these forests, goes a loss of species not found elsewhere.

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FIG. 8. Species numbers, endemics and widespread Saltatoria in % in coastal habitats near Pangani based on 61 species.

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